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Marine Pollution Bulletin 64 (2012) 2077–2082

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Marine Pollution Bulletin 64 (2012) 2077–2082 Marine Pollution Bulletin 64 (2012) 2077–2082 Contents lists available at SciVerse ScienceDirect Marine Pollution Bulletin journal homepage: www.elsevier.com/locate/marpolbul Molecular identification of green algae from the rafts based infrastructure of Porphyra yezoensis ⇑ ⇑ Qi Shen a, Hongye Li a, Yan Li b, Zongling Wang b, Jiesheng Liu a, , Weidong Yang a, a Department of Biotechnology, Jinan University, Guangzhou 510632, China b The First Institute of Oceanography, State Oceanic Administration, Qingdao 266061, China article info abstract Keywords: To provide more information on the origin of the Ulva prolifera bloom in Qingdao sea area in China from Green tide 2007 to 2011, the diversity of green algae growing on the rafts of Porphyra yezoensis on the coast in Ulva Jiangsu Province was investigated based on ITS, rbcL and 5S sequences. Eighty-four of green algal samples ITS from various sites and cruises in 2010 and 2011 were collected. According to ITS and rbcL sequences, rbcL samples from the rafts of P. yezoensis fell into four clades: Ulva linza-procera-prolifera (LPP) complex, Ulva 5S flexuosa, Blidingia sp. and Urospora spp. However, based on the 5S rDNA, a more resolved DNA marker, only one of the 84 samples belonged to U. prolifera. Combined with the previous reports, it is likely that U. prolifera bloom in Qingdao sea area might consist of more than one origin, and Porphyra cultivation rafts might be one of the causes. Ó 2012 Elsevier Ltd. All rights reserved. 1. Introduction images from 2008 to 2009, the drifted biomass initiated offshore of the coast of Jiangsu Province and was transported across the The excessive growth of green algae species such as Ulva, Enter- Yellow Sea to Qingdao coast by seasonal winds and surface cur- omorpha, Chaetomorpha and Cladophora, has been reported in the rents (Sun et al., 2008; Liu et al., 2009). However, the original formation of macroalgal blooms or green tide events in many parts ‘‘seed’’ source of the drifting bloom remained unclear. Some of the world including Europe, North America, South America, reports proposed that the accumulation and disposal of waste U. Japan and Australia (Fletcher, 1996; Morand and Briand, 1996; prolifera from Porphyra cultivation rafts was the most probable Hiraoka et al., 2004; Morand and Merceron, 2005; Merceron cause of the blooms (Keesing et al., 2011; Wang et al., 2007; Liu et al., 2007). One of the green tide algae, the filamentous alga Ulva et al., 2009). U. prolifera was the dominant fouling species growing prolifera, formerly known as Enteromorpha prolifera (Hayden et al., on the rafts based infrastructure of Porphyra yezoensis aquaculture. 2003), is broadly distributed along the nearshore coasts of the It was estimated that about 91–505 kg/ha U. prolifera was attached north-eastern Asia (Shimada et al., 2008). In the Yellow Sea of to P. yezoensis in the coast of Jiangsu Province, and a total biomass China, large-scale green algal blooms have occurred for five con- came up to 4956 tons during the harvesting of P. yezoensis (Liu secutive years from 2007 to 2011 (Jiang et al., 2008; Sun et al., et al., 2010a). This was sufficient to seed a bloom when they were 2008; Tian et al., 2011). Especially in the summer of 2008, the dislodged from the rafts as a result of harvesting practice (Liu et al., world’s largest green tide occurred along the coast of the Yellow 2010a). However, based on the ribotype analysis of the free-float- sea near Qingdao, China, which caused severe social problems as ing U. prolifera samples in 2008 and 2009 blooms, Duan et al. well as marine ecological issues. The dominant bloom algal species (2012) proposed that the bloom might be derived from the Sea in 2007–2009 was identified to be the Ulva linza-procera-prolifera of Japan. Similarly, Pang et al. (2010) found that the haplotypes (LPP) based on ITS and rbcL analysis (Hayden et al., 2003; Leliaert of the Yellow Sea free-floating U. prolifera were closely related to et al., 2009; Wang et al., 2010; Liu et al., 2010b,c). Later, Duan those from Japanese coast but less to European and American et al. (2012) amended it to U. prolifera based on 5S phylogenetic algae. Together with the similarity of samples from P. yezoensis analysis. farming rafts to U. linza in the morphology, they presented that It is crucial to correctly identify the origin of the bloom for land-based animal aquaculture ponds along the Jiangsu Province understanding the large-scale green tide and exploring solutions coast were the source of the green tide algae. to the problems it can potentially cause. According to satellite To provide more information on the origin of the massive drift- ing green tide, here we investigated the diversity of the green algae ⇑ Corresponding authors. Tel.: +86 20 85228470. growing on the rafts of P. yezoensis in the Gaoni, Niluosha, E-mail address: [email protected] (W. Yang). Xiaoyangkou of the Jiangsu Province in China from November 0025-326X/$ - see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.marpolbul.2012.07.021 2078 Q. Shen et al. / Marine Pollution Bulletin 64 (2012) 2077–2082 2010 to April 2011. The growing green algae twined on the rafts, 2.3. DNA extraction with highly variable morphology at different developmental stages and environmental conditions. So, morphological identification The fresh algal samples were washed three times with sterilized was insufficient to distinguish among the different algae species, water, and then dried with filter paper. Unialgal material for each especially among the Ulva genus. In this study, the diversity of sample was detached carefully for DNA extraction. Total DNA was the green algae was investigated based on the phylogenetic analy- extracted according to the manual of HP plant DNA extract kit sis with the sequences of nuclear encoded ribosomal DNA internal (Omega, USA). DNA quality was examined by 1% TAE agarose gels transcribed spacer region (ITS nrDNA) and the plastid encoded stained with GoldView. large subunit of ribulose-1,5-bisphosphate carboxylase/oxgenase gene (rbcL). In addition, 5S rDNA phylogenetic analysis was con- 2.4. ITS rDNA, rbcL gene and 5S rDNA spacer amplification and ducted to distinguish among the species in the LPP complex sequencing (Shimada et al., 2008; Duan et al., 2012). PCR primers for ITS, rbcL and 5S listed in Table 2 were synthe- 2. Materials and methods sized by Shanghai Sangon Biologic Engineering Technology and Service Co. Ltd., China. The PCR amplifications of ITS nrDNA and 2.1. Collection of samples rbcL genes were performed as described by Leskinen and Pamilo (1997) and Manhart (1994). As for 5S rDNA, the PCR was con- Rudong (RD), is the main coastal city for P. yezoensis aquacul- ducted as reports by Shimada et al. (2008) and Duan et al. ture in Jiangsu Province. It is located in the south-western coast (2012), in which the primer pair 5SF-5SR locates to the 5S rDNA of the Yellow Sea. So, three sites near to RD, Xiaoyangkou, Gaoni tandem arrays amplified multiple DNA fragments. Total genomic and Niluosha were chosen for exploring the distribution of green DNA (30–40 ng) was added to 50 lL PCR reactions containing 1Â algae attached to the P. yezoensis. Green algae samples at the three PCR buffer (Takara, Dalian, China), 0.8 mM dNTPs (Takara), sites (Fig. 1) were taken monthly from November-2010 to May- 25 mM of each primer and 1.6 U Taq Polymerase (Takara). PCR 2011 (Table 1). To learn the relationship between diversity of the was carried out in a MJ MiniTM Gradient Thermal Cycler (BIO- green algae attached to the P. yezoensis and dominant bloom algal RAD). PCR profiles for different genes were set as follows: ITS species in Qingdao, totally 84 samples from P. yezoensis and 3 sam- nrDNA amplification included an initial denaturation at 94 °C for ples (qingdao1, qingdao2 and qingdao 724) from Qingdao sea area 5 min, followed by 35 cycles of 94 °C for 1 min 10 s, 54 °C for in China were collected during the 2011 bloom. 50 s and 72 °C for 1 min 30 s; the rbcL gene was amplified accord- ing to the reaction profile (94 °C for 3 min, followed by 35 cycles of 94 °C for1 min, 45 °C for 2 min, and 65 °C for 3 min) and the final 2.2. Treatment of algae step at 72 °C for 10 min; the 5S amplification reaction profile in- cluded an initial denaturation at 94 °C for 5 min, followed by 35 The green algae collected from the rafts of P. yezoensis were cycles of 94 °C for 1 min, 55 °C for 45 s and 72 °C for 45 s, and a fi- cleaned in situ and brought back to the laboratory in cooled box nal extension at 72 °C for 10 min. The PCR products were resolved within 24 h. After cleaning again with sterilized seawater, the algal by 1.0% agarose gel electrophoresis, excised by the gel purification samples in good growth condition were sorted out and cultured in method using the QIAquick DNA Gel Purification Kit (QIAGEN, USA) PES medium (Berges et al., 2001) for one week to remove epiphytic and sequenced by Invitrogen (Invitrogen, China). When the ITS and diatoms for further DNA analyses. 5S rDNA spacer fragments could not be directly sequenced, PCR products were purified using the QIAquick DNA Gel Purification Shandong province Kit (QIAGEN). PCR products were cloned into pMD19-T Vector Qingdao (Takara, China) according to the manufacturer’s instructions, and sequenced by Invitrogen (Invitrogen, China).
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